Psyttalia Walker, 1860

Taxonomic History / Nomenclature
Psyttalia Walker, 1860: 311: Type species Psyttalia testacea Walker, 1860: 311 (monobasic).

Synonyms relevant to fruit-infesting tephritid parasitoids: Austroopius Szépligeti, 1900 (synonymized by Wharton 1987) and Acidoxanthopius Fischer, 1972 (synonymized by Wharton 1997).

Psyttalia was first recognized as an opiine by Muesebeck (1931), who synonymized it with Opius and renamed the type species walkeri because of secondary junior homonym in Opius. Prior to 1987, therefore, most species now in Psyttalia were placed in Opius. Psyttalia was treated as a subgenus of Opius by Fischer (1972), then elevated to generic rank by Wharton (1987). Wharton (1987) proposed Austroopius as a subgenus of Psyttalia, but later (Wharton 1997, 2009) abandoned subgenera due to lack of unambiguous features useful for defining each taxon as monophyletic. Wharton (1997) combined morphological and biological characters to suggest three informal species groups within Psyttalia, and also placed Acidoxanthopius as a synonym of Psyttalia.

Type locality of Psyttalia testacea: Sri Lanka; primary type in The Natural History Museum, London (Walker did not indicate how many specimens he had in the original description).

Valid genus (Wharton 1987, 2009)

Description
Occipital carina nearly always present laterally, extending dorsally more than half height of head (fletcheri, Fig. 1), though absent in Psyttalia insignipennis (Granger). Labrum broadly exposed beneath short, truncate or crescentic clypeus (incisi, Fig. 2 and lounsburyi, Fig. 4); clypeus usually weakly protruding, its ventral surface and junction with top of labrum clearly visible. First flagellomere equal to or, more commonly, slightly longer than second. Propleuron always without oblique carina dorsad propleural flange. Notauli often deep, but completely confined to anterior declivity of mesoscutum (thus largely absent: lounsburyi, Fig. 6); always unsculptured. Midpit absent. Postpectal carina absent. Hind tibia dorso-posteriorly without basal carina. Fore wing (incisi, Fig. 11 and concolor, Fig. 12) with second submarginal cell long to very long; m-cu arising basad or directly in line with 2RS, only rarely arising distad 2RS in species attacking fruit-infesting Tephritidae; the vein between m-cu and 2RS often distinctly thickened. Hind wing (incisi, Fig. 14) with RS absent basally, represented at most by a weak crease distally; m-cu completely absent. Metasoma of several species with second tergum finely granular or coriaceous, at least basally, metasoma otherwise unsculptured beyond petiole. Ovipositor varying from relatively short (protruding about one-third to one-half length of metasoma beyond apex) to very long. For additional details see Wharton (1987) and especially Wharton (2009).
3055_mximage
1. P. fletcheri head: late...
2968_mximage
2. P. incisi face: anterior...
2974_mximage
3. P. incisi face: anteri...
2926_mximage
4. P. lounsburyi face: an...
2924_mximage
5. P. lounsburyi face: anterior...
2928_mximage
6. P. lounsburyi thora...
2972_mximage
7. P. incisi tho...
2973_mximage
8.P. incisi mesosoma: dorsal oblique
2965_mximage
9. P. incisi propodeu...
3052_mximage
10.P. fletcheri wings
2960_mximage
11.P. incisi fore wing
3070_mximage
12.P. concolor fore wing
3056_mximage
13.P. fijiensis fore wing
2962_mximage
14.P. incisi hind wing
2976_mximage
15. P. incisi habitus: latera...
3053_mximage
16. P. fletcheri habitus: lateral...
3068_mximage
17. P. cosyrae habitus: lat...
3074_mximage
18.P. concolor habitus: lateral
2927_mximage
19. P. lounsburyi habitus: later...
2923_mximage
20. P. lounsburyi Metasoma...
2922_mximage
21. P. lounsburyi metasoma: lateral...
 
Diagnosis and Relationships
Psyttalia is distinguished from other species still included in Opius by the combination of a relatively short second metasomal tergum, short clypeus, strongly attenuate hypopygium, and absence of both hind wing m-cu and at least the basal half of hind wing RS. The shortened second tergum is one of the more important features defining this group, but is not easy to see, has not been compared across the genus Opius (in which at least a few species have second and third terga roughly equal in length), and varies within Psyttalia from species with the second tergum extremely short to those in which the second tergum is short, but roughly equal in length to the third. For purposes of recognition, useful features in addition to those listed above include a large second submarginal cell and the absence of a mesonotal midpit. The occipital carina is also well developed laterally in all but one species. The only other tephritid parasitoids with a short clypeus, a long second submarginal cell, and a well-developed occipital carina (primarily in the genus Utetes) also have a well-developed midpit on the mesoscutum.
Biology / Hosts
Host records are known for about 40% of the described species of Psyttalia , and these are restricted to the family Tephritidae. About 80% of the records are from fruit-infesting tephritids, with the remainder attacking flower-infesting tephritids. Those attacking tephritids in flower heads are, as a group, morphologically distinct from those attacking tephritids in fruits, and based on field-collected material, there is almost no overlap in host habitat. The evolution of host shifts in these groups of Psyttalia would be an interesting topic to explore.

Extensive information is available on the biology of Psyttalia concolor (Szépligeti), P. humilis (Silvestri), and P. fletcheri (Silvestri) in association with their use in biological control. More limited information is available on P. incisi (Silvestri) in this regard. Psyttalia concolor was introduced to Italy in an effort to control olive fly shortly after its discovery in Tunisia. Its early use in Italy has been well documented (Silvestri, 1922, 1938; Delucchi, 1957), as has its subsequent use in augmentation programs following development of mass rearing techniques using medfly as hosts. As a result of these efforts, there is now a considerable amount of information on the developmental biology of Psyttalia concolor, as well as other facets of its biology related to its utility for biological control of fruit pests (Féron, 1954; Ragusa, 1957; Biliotti and Delanoue, 1959; Delanoue, 1960, 1961; Arambourg, 1962; Genduso, 1967; Liotta, 1969; Raspi and Loni, 1994; Loni, 1997; Canale, 1998; Raspi and Canale, 2000; Canale and Raspi, 2000). P. concolor and P. lounsburyi have been reared from Bactrocera oleae collected from Olea europaea cuspidata in Kenya (Copeland et al. 2004). In experimental studies, P. concolor successfully parasitized second instar larvae of Ceratitis capitata (Wiedemann), and first and second instar larvae of Bactrocera oleae (Rossi) (Canale 1998, Raspi and Canale 2000). P. concolor siculus was introduced in Bolivia in 1968 to control Ceratitis capitata (Bennett and Squire 1972).

Psyttalia humilis, originally collected in South Africa, was successfully established in Hawaii in 1913 against medfly and a detailed biology was published by Pemberton and Willard (1918).

During the exploration phase of the Oriental fruit fly program, several of the opiines from Kenya were variously identified as color varieties of Psyttalia concolor or as Psyttalia perproxima (Clausen et al. 1965). Material from the same localities had been identified as either P. humilis or P. perproxima during an earlier sampling program (Bianchi and Krauss, 1936). Difficulty in identification of these three species is still a problem, and uncertainty over whether or not they are distinct makes it difficult to correctly associate previously published host records. See Rugman-Jones et al. (2009) for a more detailed treatment of this problem.

Much of the biological information on Psyttalia fletcheri and Psyttalia incisi is summarized by Clausen (1978). More recent information can be found in Ramadan et al. (1991). Other species, originating from Kenya, are currently being cultured in France, Guatemala, California, and Hawaii; see species pages for Psyttalia humilis , Psyttalia concolor , Psyttalia lounsburyi , and Psyttalia ponerophaga .

There are numerous host records for members of the expanded concolor species group as defined here. A few species, notably P. efoveolata, P. inquirenda, P. somereni, and P. walkeri, have been reared only from fruits, with the host fly unknown (Silvestri 1913; Fischer 1972a, b, c). Three of the species, P. dacicida, P. lounsburyi, and P. ponerophaga, are parasitoids of olive fly, B. oleae (Silvestri 1912, 1913, 1916b; Copeland et al. 2004; Sime et al. 2007; Daane et al. 2008), and have thus far been recorded only from this host. Psyttalia concolor is also a parasitoid of B. oleae and was originally described from specimens reared from olives. It is capable of attacking a wide variety of other fruit-infesting tephritids both in its native range and in areas where it has been introduced. In addition to Bactrocera, known hosts include tephritid species in the genera Anastrepha Schiner, Ceratitis MacLeay, Capparimyia Bezzi, Carpomya Costa, and Dacus Fabricius (Wharton and Gilstrap 1983). Psyttalia makii has been recorded from both Bactrocera and Carpomya (Wharton and Gilstrap 1983) while P. dexter, P. perproxima, and P. phaeostigma have all been reared from various species of Dacus ( SIlvestri 1913; Steck et al. 1986; Kimani-Njogu et al. 2001). Psyttalia perproxima is primarily a parasitoid of various Ceratitis and Trirhithrum Bezzi species while P. phaeostigma, which is mainly known as a parasitoid of Dacus ciliatus Loew and other cucurbit pests, has additionally been recorded from Ceratitis and Carpophthoromyia. Psyttalia cosyrae, P. distinguenda, P. humilis, and P. insignipennis have all been reared from species of Ceratitis (Silvestri 1913; Wilkinson 1927; Wharton et al. 1999; Mohamed et al. 2003, though insignipennis may have a broader host range ( Wharton et al. 1999) and humilis may have been reared on other hosts at least briefly during attempts to redistribute it from Hawaii for biological control of other tephritid pests (Clausen 1978; Wharton 1989). For a recent species-level treatment with additional host records, see Rugman-Jones et al. (2009). The temperate species P. ophthalmica and P. rhagoleticola are both parasitoids of Rhagoletis ( FIscher 1972b; Tobias 1977) and P. brevitemporalis was described from specimens reared from a species of Myoleja ( Tobias (1998)). Finally, Silvestri (1913) recorded P. inconsueta from Carpophthoromyia tritea Walker. Though Fischer (1987) placed inconsueta in his group B, based on the wing venation as illustrated by Silvestri (1913), the species is otherwise more similar in sculpture and facial features to other members of the concolor species group, and at least one of the wings on the type series has fore wing m-cu interstitial rather than postfurcal.

Although the hosts recorded above for the concolor species group are fruit-infesting tephritids, the only known host of P. dexter develops in fruits that are pod-like (Silvestri 1913). Similarly, I have seen specimens that are not easily distinguished from P. concolor, reared from Coelotrypes Bezzi infesting flowers of Convolvulaceae. Thus, a few caveats need to be attached to the generalizations about the types of hosts attacked by members of the concolor species group. Also, because of the evidence for host associated differentiation in this group, as exemplified by P. halidayi, published host records need to be carefully verified.

Distribution
Psyttalia is a fairly large, Old World genus, with about 80 described species (Wharton 2009). Maximum diversity occurs in the region from Africa east through India and Southeast Asia. One species, Psyttalia concolor, has been distributed throughout much of the world for biological control programs against various pest tephritids. Of the other species with confirmed rearings from fruit-infesting tephritid hosts, some additional comments on differentiation of species are presented here, by region, for 3 Palaearctic species, 2 species found on islands in the southwestern Indian Ocean, 7 species from the Indo-Pacific, and 3 species from the Afrotropics. Two of the Indo-Pacific species, Psyttalia incisi and Psyttalia fletcheri, are both established in Hawaii, where they were released against Oriental fruit fly, Bactrocera dorsalis (Hendel), and melon fly, Bactrocera cucurbitae (Coquillett), respectively.

Western Indian Ocean

The two species occurring in Madagascar and the Mascarenes, Psyttalia insignipennis (Granger) and Psyttalia distinguenda (Granger) are readily separated from one another by the absence of an occipital carina in insignipennis. Additional information on hosts, distribution, and recognition, based on work done largely by CIRAD, was published by Wharton et al. (1999).

Palaearctic

A key for separating two of the Palaearctic species, Psyttalia ophthalmica (Tobias) and Psyttalia rhagoleticola (Sachtleben), from Psyttalia concolor can be found in Tobias (1977). Psyttalia ophthalmica is a darker species than the other two, with a slightly longer ovipositor and a coarsely punctate face. Psyttalia rhagoleticola has distinctly longer antennae than Psyttalia concolor (approximately 40 antennal segments in Psyttalia rhagoleticola and nearly always less than 35 antennal segments in Psyttalia concolor). Psyttalia ophthalmica is known only from the Maritime Region of far eastern Russia, around Vladivostok, where it has been reared from Rhagoletis in Lonicera and briars. Psyttalia rhagoleticola has a much more western distribution, occurring from western Europe to Kazakhstan, where it attacks various species of Rhagoletis, including Palaearctic cherry fruit fly, Rhagoletis cerasi. Psyttalia rhagoleticola also attacks Myoleja lucida (Fallén) larvae (Hoffmeister 1992). The distribution of Psyttalia rhagoleticola may overlap that of the other Palaearctic species, Psyttalia ponerophaga (Silvestri), known only from a single collection in Cherat, northern Pakistan, but the latter species was reared from olive fly in apparently wild olives. Psyttalia ponerophaga is one of the few parasitoids of fruit-infesting tephritids in this genus in which the fore wing m-cu enters the second submarginal cell. It is thus readily identified relative to the other 3 species discussed in this paragraph. Psyttalia concolor has been collected from olive fly, Bactrocera oleae, in Jordan (Mustafa and Al-Zaghal 1987) and Crete (Bigler et al. 1986).

Indo-Pacific

The 7 species from the Indo-Pacific region include 2 species, Psyttalia makii (Sonan) and Psyttalia walkeri (Muesebeck), with venation somewhat similar to that found in Psyttalia concolor (concolor, Fig. 1). The vein segment between m-cu and 2RS is usually distinctly thickened in part in Psyttalia makii but barely so or not at all in Psyttalia walkeri. The base of the second metasomal segment tends to be more polished in these two species than in Psyttalia concolor. Psyttalia makii was originally described from Taiwan but was subsequently recorded from the Philippines and Thailand. Psyttalia walkeri, originally collected in Sri Lanka, has also been recorded from Indonesia and Malaysia. Both species have been reared from various hosts in the Bactrocera dorsalis complex, as well as a few other species of Bactrocera.

The fletcheri species group. Three of these species, Psyttalia fijiensis, Psyttalia muesebecki, and Psyttalia novaguineensis have fore wing 2RS thickened medially (fijiensis, Fig. 2). The three species with a medially thickened 2RS have generally been separated from one another by coloration of the fore wing and sculpture of the propodeum (Fischer 1987), but both these features are somewhat variable. Psyttalia fijiensis tends to have the fore wing darkened medially while the fore wing of the other two species tends to be more uniformely hyaline. Psyttalia muesebecki apparently lacks a median carina on the propodeum but this is present in Psyttalia novaguineensis. Psyttalia fijiensis is more widely distributed than the other two, and consequently has a wider range of hosts (Wharton and Gilstrap 1983). Psyttalia muesebecki is known only from New Caledonia.

The two remaining species, Psyttalia fletcheri and Psyttalia incisi, are very similar to one another in the curvature of fore wing m-cu and distally enlarged subdiscal cell, but m-cu is more basally displaced in Psyttalia incisi, resulting in a different pattern of thickening ( incisi, Fig. 2 and fletcheri, Fig. 1). Both species were originally described from India and subsequently introduced to and established in Hawaii. Psyttalia fletcheri attacks the melon fly, Bactrocera cucurbitae and Psyttalia incisi attacks Oriental fruit fly, Bactrocera dorsalis. Additional host and distribution records can be found in Wharton and Gilstrap (1983).

Afrotropics

Two species, Psyttalia cosyrae (Wilkinson) and Psyttalia phaeostigma (Wilkinson) have distinctly longer ovipositors (cosyrae, Fig. 6) than all of the others. Psyttalia cosyrae has been reared from Ceratitis cosyra infesting mangoes and a few wild host plants in Kenya and Tanzania. Psyttalia phaeostigma has been reared from Dacus ciliatus infesting cucurbits in Kenya. Psyttalia lounsburyi has a distinctly darker color pattern than most other Afrotropical species of Psyttalia. Considerable information on speciation in the Afrotropics can be found in Rugman-Jones et al. (2009).

3052_mximage
1.P. fletcheri wings
2960_mximage
2.P. incisi fore wing
2927_mximage
3. P. lounsburyi habitus...
3074_mximage
4. P. concolor habitus: female...
3070_mximage
5.P. concolor fore wing
3068_mximage
6. P. cosyrae habitus: fem...
3056_mximage
7.P. fijiensis fore wing
 
Distribution
No referenced distribution records have been added to the database for this OTU.
Map

There are no specimens currently determined for this OTU, or those specimens determined for this OTU are not yet mappable.

Remarks
Psyttalia is a moderately large genus. I recognize 79 valid species as of 2009 (Wharton 2009), largely following the works of Fischer (1972a, b, c, 1987, 1988, 1989, 1990, 1996, 2000). Fischer provided much of the fundamental work on Psyttalia, following its initial recognition as an opiine by Muesebeck (1931). Fischer (1963, 1972a, b, c, 1987) was the first to describe the distinctive metasomal features that separate Psyttalia from other opiines, and to note the similarities between Psyttalia, Austroopius Szépligeti and Opius (Acidoxanthopius Fischer). Fischer (1972b, c, 1987) also provided the morphological basis for delimiting species groups within Psyttalia, focusing on wing venation and propodeal sculpture.

For additional information, especially on use in biological control, see the Psyttalia page on the website devoted to parasitoids of fruit-infesting Tephritidae: http://mx.speciesfile.org/projects/8/public/public_content/show/13190?content_template_id=88

Acknowledgements
This page was assembled by Bob Wharton. It is part of a review of the genera of World Opiinae, conducted at Texas A&M University. We are particularly grateful to Xanthe Shirley, Andrew Ly, Patricia Mullins, Trent Hawkins, Lauren Ward, Cheryl Hyde, Karl Roeder, Danielle Restuccia, and Andrea Walker, who did nearly all of the imaging for this project. Matt Yoder and Istvan Miko provided guidance on databasing issues associated with our use of mx and HAO respectively. This project would not have been possible without the kindness of many curators at museums throughout the world who gave generously of their time to Bob Wharton and his students. In particular, I thank Henry Townes (deceased) and David Wahl (American Entomological Institute, Gainesville), Gordon Nishida (Bernice P. Bishop Museum, Honolulu), Norm Penny, and Bob Zuparko (California Academy of Sciences, San Francisco), Bill Mason (deceased), Mike Sharkey, Andrew Bennett, and Henri Goulet (Canadian National Collection, Ottawa), Paul Dessart (deceased) (Institut Royal des Sciences Naturelles de Belgique, Brussels), Marc De Meyer (Koninklijk Museum voor Midden-Afrika, Tervuren), Axel Bachmann (Museo Argentino de Ciencias Natureles, Buenos Aires), Eberhard Koenigsmann (deceased) and Frank Koch (Museum fuer Naturkunde der Humboldt-Universitaet, Berlin), J. Casevitz Weulersse and Claire Villemant (Museum National d’Historie Naturelle, Paris), James O’Connor (National Museum of Ireland, Dublin), Jenö Papp (National Museum of Natural History, Budapest), Kees van Achterberg (National Museum of Natural History, Leiden), Max Fischer, Herb Zettel, and Dominique Zimmermann (Naturhistorisches Museum, Wien), Per Persson and Lars-Åke Janzon (Naturhistoriska Riksmuseet, Stockholm), Ermenegildo Tremblay (Silvestri Collection, Portici), Erasmus Haeselbarth (Staatliche Naturwissenschaftliche Sammlungen Bayerns, Munich), Tom Huddleston and Gavin Broad (The Natural History Museum, London), Paul Marsh and Robert Kula (USDA Systematic Research Laboratory and US National Museum of Natural History, Washington, D. C.), Vladimir Tobias (deceased) and Sergey Belokobylskij (Zoological Institute, Academy of Sciences, St. Petersburg), and Roy Danielsson (Zoological Institute, Department of Systematics, Lund) for facilitating loans and general assistance associated with examination of holotypes and other material in their care. This work was supported largely by NSF/PEET DEB 0328922 and 0949027, with REU supplements 0616851, 0723663, 1026618, 1213790, 1313933 (to Wharton). This project was also supported by the Initiative for Future Agriculture and Food Systems grant no. 00-52103-9651 from the USDA/CSREES, USAID grant no. PCE-G-00-98-0048-00 (the latter two in collaboration with ICIPE in Kenya), and in part by a contract from the California Department of Food and Agriculture. Page last updated July, 2014. The material on this page is freely available, but should be acknowledged if used elsewhere.

This material is based upon work supported by the National Science Foundation under Grant Numbers DEB 9300517, DEB (PEET) 9712543, DEB (PEET) 0328922 with REU supplements 0723663 and 1026618 and DEB 0949027 with REU supplements 1213790 and 1313933. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.